Visual Standards and Disciplinary Change: Normal Plates,Tables and Stages in Embryology

Maniatis T. Fritsch E. F. Sambrook J. , Molecular cloning: A laboratory manual (Cold Spring Harbor, N.Y., 1982); Nieuwkoop P. D. Faber Job (ed.), Normal table of Xenopus laevis (Daudin): A systematical and chronological survey of the development from the fertilized egg till the end of metamorphosis, 2nd edn (Amsterdam, 1967); Gurdon John B. Hopwood Nick , “The introduction of Xenopus laevis into developmental biology: Of empire, pregnancy testing and ribosomal genes”, International journal of developmental biology, xliv (2000), 43–50.

Latour Bruno , Science in action: How to follow scientists and engineers through society (Cambridge, Mass., 1987), 250–4. On metrology, see especially O'Connell Joseph , “Metrology: The creation of universality by the circulation of particulars”, Social studies of science, xxiii (1993), 129–73; Wise M. Norton (ed.), The values of precision (Princeton, 1995); Schaffer Simon , “Metrology, metrication, and Victorian values”, in Lightman Bernard (ed.), Victorian science in context (Chicago, 1997), 438–74; and (for literature reviews) Hessenbruch Arne , “Metrology” and “Standardization”, in idem (ed.), Reader's guide to the history of science (London, 2000), 477–80 and 704–6; and on biological and medical standards, Köhler Robert E. , Lords of the fly: Drosophila genetics and the experimental life (Chicago, 1994); Hess Volker (ed.), Normierung der Gesundheit: Messende Verfahren der Medizin als kulturelle Praktik um 1900 (Husum, 1997); Sohn Werner Mehrtens Herbert (ed.), Normalität und Abweichung: Studien zur Theorie und Geschichte der Normalisierungsgesellschaft (Opladen, 1999); Zimmerman Andrew , Anthropology and antihumanism in Imperial Germany (Chicago, 2001), 86–107; and Sinding Christiane , “Making the unit of insulin: Standards, clinical work, and industry, 1920–1925”, Bulletin of the history of medicine, lxxvi (2002), 231–70.

Hopwood Nick , “Embryology”, in Bowler Peter J. Pickstone John V. (ed.), The Cambridge history of science, vi: The modern biological and earth sciences (Cambridge, forthcoming).

Daston Lorraine Galison Peter , “The image of objectivity”, Representations, xl (1992), 81–128; Galison Peter , “Judgment against objectivity”, in Jones Caroline A. Galison Peter (ed.), Picturing science, producing art (New York, 1998), 327–59; Daston Lorraine , “Objectivity versus truth”, in Bödeker Hans Erich Reill Peter Hanns Schlumbohm Jürgen (ed.), Wissenschaft als kulturelle Praxis 1750–1900 (Göttingen, 1999), 17–32.

See, e.g., Hopwood Nick , “‘Giving body’ to embryos: Modeling, mechanism, and the microtome in late nineteenth-century anatomy”, Isis, xc (1999), 462–96, pp. 490–1. Embryological experts were exposed to few of the public challenges, which, it has been suggested, elicit ‘mechanical objectivity’ as a defence; see Porter Theodore M. , Trust in numbers: The pursuit of objectivity in science and public life (Princeton, 1995); and Alder Ken , “Making things the same: Representation, tolerance and the end of the ancien régime in France”, Social studies of science, xxviii (1998), 499–545.

Wise M. Norton , “Precision: Agent of unity and product of agreement”, in idem (ed.), Values of precision (ref. 2), 92–100, 222–36 and 352–61; Schaffer Simon , “Accurate measurement is an English science”, ibid., 135–72, pp. 135–7.

For ‘ways of working’ in embryology, see Hopwood , “Embryology” (ref. 3); and in general, Pickstone John V. , Ways of knowing: A new history of science, technology and medicine (Manchester, 2000). The approach is closer to Rudwick Martin J. S. , “The emergence of a visual language for geological science, 1760–1840”, History of science, xiv (1976), 149–95.

Hopwood Nick , “Producing development: The anatomy of human embryos and the norms of Wilhelm His”, Bulletin of the history of medicine, lxxiv (2000), 29–79; idem, “Embryonen ‘auf dem Altar der Wissenschaft zu opfern’: Entwicklungsreihen im späten neunzehnten Jahrhundert”, in Duden Barbara Schlumbohm Jürgen Veit Patrice (ed.), Geschichte des Ungeborenen: Zur Erfahrungs- und Wissenschaftsgeschichte der Schwangerschaft, 17.–20. Jahrhundert (Göttingen, 2002), 237–72; idem, Embryos in wax: Models from the Ziegler studio, with a reprint of “Embryological wax models” by Friedrich Ziegler (Cambridge and Bern, 2002); idem, “Plastic publishing in embryology”, in de Chadarevian Soraya Hopwood Nick (ed.), Models: The third dimension of science (Stanford, 2004), 170–206. On developmental series in other sciences, see, e.g., Rudwick Martin J. S. , “Charles Lyell's dream of a statistical palaeontology”, Palaeontology, xxi (1978), 225–44; idem, Scenes from deep time: Early pictorial representations of the prehistoric world (Chicago, 1992); van Keuren David K. , “Museums and ideology: Augustus Pitt-Rivers, anthropological museums, and social change in later Victorian Britain”, Victorian studies, xxviii (1984), 171–89; and Armstrong David , “The temporal body”, in Cooter Roger Pickstone John (ed.), Medicine in the twentieth century (Amsterdam, 2000), 247–59.

Roe Shirley A. , Matter, life, and generation: Eighteenth-century embryology and the Haller—Wolff debate (Cambridge, 1981); Jordanova L. J. , “Gender, generation and science: William Hunter's obstetrical atlas”, in Bynum W. F. Porter Roy (ed.), William Hunter and the eighteenth-century medical world (Cambridge, 1985), 385–412; Hagner Michael , “Enlightened monsters”, in Clark William Golinski Jan Schaffer Simon (ed.), The sciences in enlightened Europe (Chicago, 1999), 175–217; Duden Barbara , “Zwischen ‘wahrem Wissen’ und Prophetie: Konzeptionen des Ungeborenen”, in idem et al. (ed.), Geschichte des Ungeborenen (ref. 8), 11–48; Filippini Nadia Maria , “Die ‘erste Geburt’: Eine neue Vorstellung vom Fötus und vom Mutterleib (Italien, 18. Jahrhundert)”, ibid., 99–127; Enke Ulrike , “Von der Schönheit der Embryonen: Samuel Thomas Soemmerrings Werk Icones embryonum humanorum (1799)”, ibid., 205–35; Wellmann Janina , “Wie das Formlose Formen schafft: Bilder in der Haller-Wolff-Debatte und die Anfänge der Embryologie um 1800”, Bildwelten des Wissens, i (2003), part 2, 105–15.

The previous histories introduce stage series. The most extensive concentrate on human embryos and draw heavily one from another: George W. Corner, preface to Streeter George L. , Developmental horizons in human embryos: Age groups XI to XXIII. Collected papers from the Contributions to embryology published by the Carnegie Institution of Washington (Washington, D.C., 1951), pp. iii–iv; O'Rahilly Ronan , Developmental stages in human embryos, including a survey of the Carnegie collection. Part A: Embryos of the first three weeks (stages 1 to 9) (Washington, D.C., 1973), 1–8; idem, “One hundred years of human embryology”, Issues and reviews in teratology, iv (1988), 81–128; idem and Müller Fabiola , Developmental stages in human embryos, including a revision of Streeter's “horizons” and a survey of the Carnegie collection ([Washington, D.C.], 1987), 1–8. Corner reviewed the shift in human embryology from the seriations of Wilhelm His and Franz Keibel to the stage systems of his own predecessors as director of the Carnegie Department of Embryology, and cited Ross Harrison's normal stages of Amblystoma as exemplary. It is not clear to me why Corner also singled out Balfour F. M. , A monograph on the development of elasmobranch fishes (London, 1878) as pioneering “the method of formal stages”.

Gould Stephen Jay , Ontogeny and phylogeny (Cambridge, Mass., 1977), 174. For revisionist views of the ‘revolt’, see Maienschein Jane , Transforming traditions in American biology, 1880–1915 (Baltimore, 1991); idem, “The origins of Entwicklungsmechanik”, in Gilbert Scott F. (ed.), A conceptual history of modern embryology (Baltimore, 1994), 43–61; Nyhart Lynn K. , Biology takes form: Animal morphology and the German universities, 1800–1900 (Chicago, 1995), 243–361; and Hopwood , “Embryology” (ref. 3).

Gould , Ontogeny and phylogeny (ref. 11); Nyhart , Biology takes form (ref. 11).

For reviews, see Churchill Frederick B. , “The rise of classical descriptive embryology”, in Gilbert (ed.), Conceptual history (ref. 11), 1–29; and Hopwood , “Embryology” (ref. 3). On morphology's institutional situation, see Nyhart , Biology takes form (ref. 11); and on the courses, Hopwood , Embryos in wax (ref. 8), 33–39.

Russell E. S. , Form and function: A contribution to the history of animal morphology (London, 1916); Gould , Ontogeny and phylogeny (ref. 11), 33–68; Appel Toby A. , The Cuvier-Geoffroy debate: French biology in the decades before Darwin (New York, 1987); Clarke Edwin Jacyna L. S. , Nineteenth-century origins of neuroscientific concepts (Berkeley, 1987), 29–100; Lenoir Timothy , The strategy of life: Teleology and mechanics in nineteenth-century German biology (Chicago, 1989), 54–111; Richards Robert J. , The meaning of evolution: The morphological construction and ideological reconstruction of Darwin's theory (Chicago, 1992), 17–61.

See the influential surveys, Valentin G. , Handbuch der Entwickelungsgeschichte des Menschen mit vergleichender Rücksicht der Entwickelung der Säugethiere und Vögel; nach fremden und eigenen Beobachtungen (Berlin, 1835); Bischoff Th. L. W. , Entwickelungsgeschichte der Säugethiere und des Menschen (Leipzig, 1842); and Kölliker Albert , Entwicklungsgeschichte des Menschen und der höheren Thiere (Leipzig, 1861).

Haeckel Ernst , Generelle Morphologie der Organismen: Allgemeine Grundzüge der organischen Formen-Wissenschaft, mechanisch begründet durch die von Charles Darwin reformirte Descendenz-Theorie (2 vols, Berlin, 1866; facsimile edn, Berlin, 1988), i, 54–55 (ontogeny), 60 (phylogeny); ii, 7, 300 (recapitulation); idem, Natürliche Schöpfungsgeschichte: Gemeinverständliche wissenschaftliche Vorträge über die Entwickelungslehre im Allgemeinen und diejenige von Darwin, Goethe und Lamarck im Besonderen, über die Anwendung derselben auf den Ursprung des Menschen und andere damit zusammenhängende Grundfragen der Naturwissenschaft, 2nd edn (Berlin, 1870), 276, 361 (biogenetic law). On concepts of recapitulation and evolution, see Gould , Ontogeny and phylogeny (ref. 11), 69–114; Churchill Frederick B. , “Weismann, hydromedusae and the biogenetic imperative: A reconsideration”, in Horder T. J. Witkowski J. A. Wylie C. C. (ed.), A history of embryology (Cambridge, 1985), 7–33; Richards , Meaning of evolution (ref. 14); and Mengal Paul (ed.), Histoire du concept de la récapitulation: Ontogenèse et phylogenèse en biologie et sciences humaines (Paris, 1993).

Groeben Christiane Müller Irmgard , The Naples Zoological Station at the time of Anton Dohrn, transl. by Richard Ivell Christl (Paris, 1975); Semon Richard , Im australischen Busch und an den Küsten des Korallenmeeres: Reiseerlebnisse und Beobachtungen eines Naturforschers in Australien, Neu-Guinea und den Molukken (Leipzig, 1896).

Haeckel Ernst , “Die Gastrula und die Eifurchung der Tiere”, Jenaische Zeitschrift für Naturwissenschaft, ix (1875), 402–508, pp. 409–12. For the confused Greek etymology of Cenogenese or Caenogenese, see Keibel Franz , “Das biogenetische Grundgesetz und die Cenogenese”, Ergebnisse der Anatomie und Entwickelungsgeschichte, vii (1898), 722–92, p. 731.

Virchow Rudolf , Die Cellularpathologie in ihrer Begründung auf physiologische und pathologische Gewebelehre: Zwanzig Vorlesungen gehalten während der Monate Februar, März und April 1858 im pathologischen Institut zu Berlin (Berlin, 1858; facsimile edn, Hildesheim, 1966), 57; Haeckel Ernst , Anthropogenie oder Entwickelungsgeschichte des Menschen: Gemeinverständliche wissenschaftliche Vorträge über die Grundzüge der menschlichen Keimes- und Stammes-Geschichte (Leipzig, 1874), 634, 717; idem, “Gastrula” (ref. 18), 412–16. Haeckel's possible borrowing from pathology is suggested by quotations from a translation of a later handbook in the Oxford English dictionary, but I am not aware that attention has been drawn to it before.

Gould , Ontogeny and phylogeny (ref. 11), 167–206.

Gegenbaur C. , “Die Stellung und Bedeutung der Morphologie”, Morphologisches Jahrbuch, i (1875), 1–19, pp. 14, 17; idem, “Cünogenese”, Verhandlungen der Anatomischen Gesellschaft, 1888, 3–9; idem, “Ontogenie und Anatomie, in ihren Wechselbeziehungen betrachtet”, Morphologisches Jahrbuch, xv (1889), 1–9.

Nyhart , Biology takes form (ref. 11), 243–77.

Hopwood , “‘Giving body’” (ref. 5).

Churchill Frederick B. , “Chabry, Roux, and the experimental method in nineteenth-century embryology”, in Giere Ronald N. Westfall Richard S. (ed.), Foundations of scientific method: The nineteenth century (Bloomington, Ind., 1973), 161–205; Maienschein , “Origins of Entwicklungsmechanik” (ref. 11); Mocek Reinhard , Die werdende Form: Eine Geschichte der Kausalen Morphologie (Marburg, 1998).

Müller Irmgard , “Die Wandlung embryologischer Forschung von der deskriptiven zur experimentellen Phase unter dem Einfluss der Zoologischen Station in Neapel”, Medizinhistorisches Journal, x (1975), 191–218; Werdinger Jeffrey , “Embryology at Woods Hole: The emergence of a new American biology”, Ph.D. dissertation, Indiana University, 1980; Pauly Philip J. , “Summer resort and scientific discipline: Woods Hole and the structure of American biology, 1882–1925”, in Rainger Ronald Benson Keith R. Maienschein Jane (ed.), The American development of biology (New Brunswick, N.J., 1991), 121–50; Harwood Jonathan , Styles of scientific thought: The German genetics community, 1900–1933 (Chicago, 1993), 30–31; Nyhart , Biology takes form (ref. 11), 243–361; Sucker Ulrich , Das Kaiser-Wilhelm-Institut für Biologie: Seine Gründungsgeschichte, seine problemgeschichtlichen und wissenschaftstheoretischen Voraussetzungen (1911–1916) (Stuttgart, 2002).

Much of the revisionist work is on the United States; on Germany, see Maienschein , “Origins of Entwicklungsmechanik” (ref. 11); Weindling Paul Julian , Darwinism and Social Darwinism in Imperial Germany: The contribution of the cell biologist Oscar Hertwig (1849–1922) (Stuttgart, 1991); Nyhart , Biology takes form (ref. 11), 243–361; Hopwood , “‘Giving body’” (ref. 5); idem, “Producing development” (ref. 8); and idem, Embryos in wax (ref. 8).

E.g., [Gabriel Gustav] Valentin , “Foetus”, in Busch D. W. H. (ed.), Encyclopädisches Wörterbuch der medicinischen Wissenschaften, xii (Berlin, 1835), 355–89, p. 355, with reference to “descriptive anatomy”; Haeckel Ernst , Generelle Morphologie (ref. 16), ii, 16; idem, Anthropogenie oder Entwickelungsgeschichte des Menschen: Keimes- und Stammes-Geschichte, 4th edn (2 vols, Leipzig, 1891), i, pp. xx–xxi; Roux Wilhelm , Correns C. Fischel Alfred Küster E. , Terminologie der Entwicklungsmechanik der Tiere und Pflanzen: Eine Ergänzung zu den Wörterbüchern der Biologie, Zoologie und Medizin sowie zu den Lehr- und Handbüchern der Entwicklungsgeschichte, allgemeinen Biologie und Physiologie (Leipzig, 1912), 91–92. The term was not only negative; a sympathetic obituarist praised Keibel's normal plates as having laid a foundation for “deskriptive Embryologie”; see Elze Curt , “Franz Keibel t. Geb. 6. Juli 1861, gest. 27. April 1929”, Klinische Wochenschrift, viii (1929), 1335.

On His, see Hopwood , “‘Giving body’” (ref. 5); and idem, “Producing development” (ref. 8); for the range of vertebrate embryology, Hertwig Oskar (ed.), Handbuch der vergleichenden und experimentellen Entwickelungslehre der Wirbeltiere (3 vols, Jena, 1906); and on the mass of new evolutionary research around 1900, Bowler Peter J. , Life's splendid drama: Evolutionary biology and the reconstruction of life's ancestry, 1860–1940 (Chicago, 1996).

His Wilhelm , “Die Entwicklung der zoologischen Station in Neapel und das wachsende Bedürfnis nach wissenschaftlichen Zentralanstalten”, Biologisches Centralblatt, vi (1886), 545–54, p. 554. All translations are mine.

Ibid., 549–54.

Gould Stephen Jay , The mismeasure of man (Harmondsworth, 1984); idem, “Ladders and cones: Constraining evolution by canonical icons”, in Silvers Robert B. (ed.), Hidden histories of science (London, 1997), 37–67. On evolutionary trees, see further especially Uschmann G. , “Zur Geschichte der Stammbaum-Darstellungen”, in Gersch Manfred (ed.), Gesammelte Vorträge übermoderne Probleme der Abstammungslehre, ii (Jena, 1967), 9–30; (with care) Bouquet Mary , “Family trees and their affinities: The visual imperative of the genealogical diagram”, Journal of the Royal Anthropological Institute, n. s., ii (1996), 43–66; and Voss Julia , Darwins Diagramme — Bilder von der Entdeckung der Unordnung (Max-Planck-Institut für Wissenschaftsgeschichte Preprint 249; Berlin, 2003); and on craniometry, Zimmerman , Anthropology and antihumanism (ref. 2), 86–107; and Hagner Michael , Geniale Gehirne: Zur Geschichte der Elitegehirnforschung (Göttingen, 2004).

von Baer Karl Ernst , Über Entwickelungsgeschichte der Thiere: Beobachtung und Reflexion, part 1 (Königsberg, 1828; facsimile edn, Brussels, 1967), 147–8.

Ibid., 4–7. On the sleepless nights, see idem, Autobiography of Dr Karl Ernst von Baer, 2nd edn, transl. by Schneider H. , ed. by Oppenheimer Jane M. (Canton, Mass., 1986), 212.

Pander Christian , Beiträge zur Entwickelungsgeschichte des Hühnchens im Eye (Würzburg, 1817), 30; idem, “Dissertation inaugurale établissant l'histoire de la métamorphose que subit l'œEuf au cours des cinq premiers jours d'incubation” [Latin original, 1817], in Les textes embryologiques de Christian Heinrich Pander (1794–1865), ed. and transl. by Schmitt Stéphane (Turnhout, 2003), 61–94.

Pander , Beiträge (ref. 34); Von Baer , Entwickelungsgeschichte (ref. 32). So much remains to be done on the visual language of embryology that even the illustrations in these major works have yet to receive serious historical attention.

Galison Daston , “Image of objectivity” (ref. 4), 84–98; His Wilhelm , Untersuchungen über die erste Anlage des Wirbelthierleibes: Die erste Entwickelung des Hühnchens im Ei (Leipzig, 1868), 56–60.

Soemmerring Samuel Thomas , “Icones embryonum humanorum”, in Schriften zur Embryologie und Teratologie, transl. by Moog Ferdinand Peter , ed. by Enke Ulrike (Basel, 2000[Latin original, 1799]), 165–89; Duden Barbara , “The fetus on the ‘farther shore’: Toward a history of the unborn”, in Morgan Lynn M. Michaels Meredith W. (ed.), Fetal subjects, feminist positions (Philadelphia, 1999), 13–25; Enke , “Schönheit der Embryonen” (ref. 9).

For a review, see Valentin , “Foetus” (ref. 27). Tables are included in Senff Carl Friedrich , Nonnulla de incremento ossium embryonum in primis graviditatis mensibus: Dissertatio inauguralis … (Halle, 1801); and Nicolai Johann August Heinrich , Beschreibung der Knochen des menschlichen Foetus: Ein Beitrag zur Anatomie des Foetus und zur Bestimmung des Alters der Embryonen und des Foetus aus der Beschaffenheit der Knochen (Münster, 1829); plates in Coste J. J. V. , Histoire générale et particulière du développement des corps organisés (Paris, 1849); and Ecker Alexander , Icones physiologicae: Erläuterungstafeln zur Physiologie und Entwickelungsgeschichte (Leipzig, 1851–9); and woodcuts in Kölliker , Entwicklungsgeschichte (ref. 15).

Von Baer , Entwickelungsgeschichte (ref. 32), pl. III.

The most striking comparisons I have found are ibid., part 2 (Königsberg, 1837; facsimile edn, Brussels, 1967), pl. IV (legend in ibid., Schlussheft, ed. by Stieda Ludwig (Königsberg, 1888), 396–7); and Wagner Rudolph , Icones physiologicae: Tabulae physiologiam et geneseos historiam illustrantes. Erläuterungstafeln zur Physiologie und Entwickelungsgeschichte (Leipzig, 1839), pl. V.

Kölliker , Entwicklungsgeschichte (ref. 15); Hertwig Oscar , Lehrbuch der Entwicklungsgeschichte des Menschen und der Wirbelthiere (Jena, 1888). Hopwood , Embryos in wax (ref. 8), 38, mislabels a figure of Hertwig's as a wood-engraving.

Haeckel , Natürliche Schöpfungsgeschichte (ref. 16), pls II–III; idem, Anthropogenie, 4th edn (ref. 27), i, pls VII-IX. Compare the wood-engravings in Balfour Francis M. , A treatise on comparative embryology, ii (London, 1881).

Gursch Reinhard , Die Illustrationen Ernst Haeckels zur Abstammungs- und Entwicklungsgeschichte: Diskussion im wissenschaftlichen und nichtwissenschaftlichen Schrifttum (Frankfurt am Main, 1981).

Daston , “Objectivity versus truth” (ref. 4), 28–29.

I am preparing a book on Haeckel's pictures of embryos.

His Wilhelm , Über die Aufgaben und Zielpunkte der wissenschaftlichen Anatomie: Rede, gehalten beim Antritt der anatomischen Professur der Universität Leipzig den 4. November 1872 (Leipzig, 1872), 9–10. Is it significant that His referred to norms (for the first time in print?) on joining the Leipzig medical faculty, where Carl Wunderlich was promoting a view of disease as deviation from a thermometrically measured norm of temperature ( Hess Volker , Der wohltemperierte Mensch: Wissenschaft und Alltag des Fiebermessens (1850–1900) (Frankfurt am Main, 2000), 146–65)?.

Hopwood , “Producing development” (ref. 8).

His Wilhelm , Anatomie menschlicher Embryonen (3 vols, Leipzig), i: Embryonen des ersten Monats (1880), 147–68 (quote on p. 147); ii: Gestalt- und Grössenentwicklung bis zum Schluss des 2. Monats (1882), 2 (quote), 23–71; iii: Zur Geschichte der Organe (1885), 236–42; Hopwood , “Producing development” (ref. 8). Use of the term ‘norm’ was not new; Gustav Valentin had written that a certain embryo could be taken “[a]s norm of the fifth week” ( Valentin , “Foetus” (ref. 27), 363), and incidentally, also of Soemmerring's “normal figures” (ibid., 357–8).

Hopwood , “Producing development” (ref. 8), 70–76.

Oppel Albert , Vergleichung des Entwicklungsgrades der Organe zu verschiedenen Entwicklungszeiten bei Wirbeltieren (Jena, 1891). There are parallels to recent concerns about gene expression data. Oppel, who was later a colleague of Roux in Halle, is perhaps more widely known as coauthor of a microscopy manual; see Eisler P. , “Albert Oppel †”, Anatomischer Anzeiger, xlviii (1915), 414–15; and Roux Wilhelm , † Oppel Albert , Archiv für Entwicklungsmechanik der Organismen, xlii (1917), 261–6.

Oppel Albert , “Dritter Theil: Entwicklungsgeschichte. Zweiter Abtheilung: Entwicklungsgeschichte der Wirbelthiere”, Jahresbericht über die Fortschritte der Anatomie und Physiologie, xx (1892), section, 608–747, pp. 683–6.

Peter Karl , “Franz Keibel: Ein Nachruf”, Anatomischer Anzeiger, lxviii (1929), 201–20; [ Fick Rudolf ], “Gedächtnisrede auf Franz Keibel”, Sitzungsberichte der Preussischen Akademie der Wissenschaften, Physikalisch-mathematische Klasse, 1929, pp. cvii–cxxii; Elze , “Keibel” (ref. 27); Stieve H. , “Franz Keibel zum Gedächtnis”, Zeitschrift für mikroskopisch-anatomische Forschung, xviii (1929), 1–4; S[treeter] G[eorge] L. , “ Keibel Franz ”, Science, lxix (1929), 637; Nauck E. Th. , Franz Keibel: Zugleich eine Untersuchung über das Problem des wissenschaftlichen Nachwuchses (Jena, 1937). Nauck celebrated Keibel as a pioneer of a holistic embryology. For an autobiographical account from 1892, see Universitätsarchiv Freiburg: B37/535.

Nyhart , Biology takes form (ref. 11), 207–40. On the Freiburg institute, see further Hopwood , Embryos in wax (ref. 8) and literature cited therein.

On Schwalbe, more prominent in histories of embryology for supervising Roux's doctoral dissertation, see Keibel Franz , “Gustav Albert Schwalbe †”, Anatomischer Anzeiger, xlix (1916), 210–21; and Le Minor J. M. Kahn J. L. , “Histoire de l'anatomie à Strasbourg”, Archives d'anatomie, d'histologie et d'embryologie normales et expérimentales, lxxii (1989), 125–55, pp. 142–7.

The excursions are mentioned as regular events in Robert Wiedersheim to Max Fürbringer, 7 June 1910, and Ernst Gaupp to Fürbringer, 18 June 1911, Universitätsbibliothek Johann Christian Senckenberg, Frankfurt am Main: Fürbringer Papers.

Keibel Franz , “Studien zur Entwicklungsgeschichte des Schweines (Sus scrofa domesticus)”, Morphologische Arbeiten, iii (1893), 1–139, pp. 1–10. The corresponding models are illustrated in Hopwood , Embryos in wax (ref. 8), 111 and 149, and their making discussed on pp. 57–59.

Keibel Franz , “Studien zur Entwicklungsgeschichte des Schweines. (Sus scrofa domesticus.) II”, Morphologische Arbeiten, v (1895), 17–168, pp. 75–78.

Idem, “Das biogenetische Grundgesetz” (ref. 18), 731–6.

Keibel Franz , “Normentafeln zur Entwickelungsgeschichte der Wirbeltiere”, Anatomischer Anzeiger, xi (1895), 225–34; idem, “Mitteilungen über die ‘Normentafeln zur Entwickelungsgeschichte der Wirbeltiere’”, ibid., xi (1896), 593–6. Personal approaches include Keibel to Max Fürbringer, 17 and 24 October 1895, Fürbringer Papers (ref. 55), the latter mentioning Keibel Semon to Merkel Friedrich , 17 October 1895, Niedersächsische Staats- und Universitätsbibliothek Göttingen: Philos. 187, 14; and Keibel to Reighard Jacob , 17 October 1895 and 15 June 1897, Bentley Historical Library, University of Michigan: Reighard Papers, box 1; none of these led directly to a publication in the series, but see Reighard Jacob , “Exhibition of figures for a Normentafel of Amia with an account of methods of photographing the embryo”, Science, xi (1900), 251. Keibel also presented the project to the German Anatomical Society and received His's support (Verhandlungen der Anatomischen Gesellschaft, 1896, 81); see also His Wilhelm , “Ueber wissenschaftliche Centralanstalten und speciell über Centralanstalten zur Förderung der Gehirnkenntniss”, Berichte über die Verhandlungen der mathematisch-physischen Classe der Königlich Sächsischen Gesellschaft der Wissenschaften zu Leipzig, liii (1901), 413–36, p. 427. Keibel surely used conferences to recruit internationally too. On serial publication, see, e.g., Howsam Leslie , “Sustained literary ventures: The series in Victorian book publishing”, Publishing history, xxxi (1992), 5–26.

Keibel , “Entwicklungsgeschichte des Schweines” (ref. 56), 5; idem, “Ueber den Entwickelungsgrad der Organe in den verschiedenen Stadien der embryonalen Entwickelung der Wirbeltiere”, in Handbuch der vergleichenden und experimentellen Entwicklungslehre der Wirbeltiere, ed. by Hertwig Oskar , iii/3 (Jena, 1906), 131–48. For a contemporary history, see Rádl Emanuel , “Ueber die Bedeutung des Prinzips von der Korrelation in der Biologie”, Biologisches Centralblatt, xxi (1901), 401–16, 490–6, 550–60, 585–91, 605–21; and for an experimentalist's statement, Spemann Hans , “Zum Problem der Correlation in der tierischen Entwicklung”, Verhandlungen der Deutschen Zoologischen Gesellschaft, xvii (1907), 22–48. In work on the statistical norming of development, Alexander Gurwitsch took a distinctive approach to this complex of problems; see Gurwitsch Alexander , “Über Determination, Normierung und Zufall in der Ontogenese”, Archiv für Entwicklungsmechanik der Organismen, xxx (1910), 133–93; and Beloussov Lev V. Opitz John M. Gilbert Scott F. , “Life of Alexander G. Gurwitsch and his relevant contribution to the theory of morphogenetic fields”, International journal of developmental biology, xli (1997), 771–9.

Keibel occasionally encouraged students to pursue developmental mechanical investigations, but — As one might expect of a Hertwig student ( Weindling , Darwinism (ref. 26), 119–30) — Was critical of Roux's inaccuracies and exaggerations; see Keibel Franz , “Bemerkungen zu Roux's Aufsatz: ‘Das Nichtnötigsein der Schwerkraft für die Entwickelung des Froscheies’”, Anatomischer Anzeiger, xxi (1902), 581–91; idem, “Bemerkung zu Wilhelm Rouxs Aufsatz: ‘Über die Ursachen der Bestimmung der Hauptrichtungen des Embryo im Froschei’”, ibid., xxiii (1903), 224. Keibel's most extensive reflections on the subject include this statement: “We have, then, every reason to welcome experimental embryology and can readily overlook the way that some of its representatives too optimistically believe themselves already to be near to generating living beings in the test-tube” ( Keibel Franz , Über experimentelle Entwicklungsgeschichte: Rede, gehalten am 27. Januar 1917 zur Feier des Geburtstages Sr. Majestät des Kaisers in der Aula der Kaiser Wilhelms-Universität Straßburg (Strasbourg, 1917), 26). Keibel greatly admired Harrison Ross : “His works are of the very greatest importance; I wish we had such a good man in Germany” ( Keibel to Mall Franklin P. , 16 December 1907, Alan Mason Chesney Medical Archives, Johns Hopkins Medical Institutions, Baltimore: Carnegie Institution of Washington Department of Embryology Papers, record group 1, series 1, box 10, folder 33).

Keibel , “Das biogenetische Grundgesetz” (ref. 18), 758 (emphasis here and elsewhere in the originals), reviewing idem, “Entwicklungsgeschichte des Schweines. II” (ref. 57), 18. Keibel derived this stance from a review by Roux's former colleague and his own predecessor as a prosector in Freiburg, the Swiss anatomist Hans Strasser.

NT 1 (i.e., the first volume in Keibel's series; see Table 1), 1–9.

His , Aufgaben und Zielpunkte (ref. 46), 9.

Schwalbe G. , “Über Variation”, Verhandlungen der Anatomischen Gesellschaft, xii (1898), 2–15. The classic work is Bateson William , Materials for the study of variation treated with especial regard to discontinuity in the origin of species (London, 1894). For (Berlin) anthropologists' caution towards Darwinism, see Zimmerman , Anthropology and antihumanism (ref. 2).

Schwalbe G. Pfitzner W. , “Varietätenstatistik und Anthropologie”, Anatomischer Anzeiger, iv (1889), 705–14; Pfitzner W. , “Beiträge zur Kenntniss des menschlichen Extremitätenskelets: Erster Beitrag”, Morphologische Arbeiten, i (1891), 1–17, p. 6 (quote); Mehnert Ernst , “Bericht über die Leichenmessungen am Strassburger anatomischen Institute”, ibid., iv (1894), 1–29. On Mehnert and Pfitzner, see Schwalbe G. , “Wilhelm Pfitzner †”, Zeitschrift für Morphologie und Anthropologie, v, no. 3 (1903), pp. v–xii, also in Anatomischer Anzeiger, xxii (1903), 481–7; and idem, “Ernst Mehnert †”, ibid., 387–92.

Mehnert Ernst , “Die individuelle Variation des Wirbelthierembryo: Eine Zusammenstellung”, Morphologische Arbeiten, v (1895), 386–444, pp. 399, 430.

Gould , Ontogeny and phylogeny (ref. 11), 174–5.

NT 1, 80–81 (quote); Mehnert Ernst , “Allgemeine Entwicklungsgeschichte: II. Variation, Heredität. A. Variation”, Jahresberichte über die Fortschritte der Anatomie und Entwicklungsgeschichte, n. s., iii (1898), 327–36, pp. 331–2; Keibel Franz , “Das biogenetische Grundgesetz” (ref. 18), 759–68; Mehnert , “Bemerkungen zu Keibels Kritiken und Referaten”, Anatomische Hefte, xii (1899), 549–65; Keibel , “Zu Mehnerts Bemerkungen über meine Kritiken und Referate”, ibid., 567–73; Mehnert , “K. E. v. Baer als Begründer der Erkenntnis der individuellen Variation im Embryonalleben”, Biologisches Centralblatt, xix (1899), 443–55; Keibel , “Bemerkungen zu Mehnert's Aufsatz: K. E. v. Baer als Begründer der Erkenntnis der individuellen Variation im Embryonalleben”, ibid., 744–6; NT 2, 13.

Mehnert Ernst , “Kainogenese: Eine gesetzmässige Abänderung der embryonalen Entfaltung in Folge von erblicher Uebertragung in der Phylogenese erworbener Eigenthümlichkeiten. Eine biologische Studie”, Morphologische Arbeiten, vii (1897), 1–156; idem, Biomechanik erschlossen aus dem Principe der Organogenese (Jena, 1898); Keibel , “Das biogenetische Grundgesetz” (ref. 18), 776–81, 787–91. Grateful by this time for any specialist support, Haeckel praised Mehnert ( Haeckel , Anthropogenic oder Entwickelungsgeschichte des Menschen: Keimes- und Stammesgeschichte, 5th edn (2 vols, Leipzig, 1903), i, 12–13), but accused Keibel, although “he has himself in very careful descriptive-embryological works provided a great number of supports for the Biogenetic Law”, of failing to understand it or to grasp “even the important distinction between palingenesis and cenogenesis” ( Haeckel , Die Lebenswunder: Gemeinverständliche Studien über Biologische Philosophie. Ergänzungsband zu dem Buche über die Welträthsel (Stuttgart, 1904), 439). Haeckel's secretary reckoned Keibel “one of the most zealous opponents of the Biogenetic Law”. “In his embryological investigations he sees only the cenogenetic alteration of the germinal form; and when contemplating the B. L. he has eyes only for undisturbed palingenesis.” There was nothing new about the extreme cenogenesis of mammalian development; Haeckel himself had pointed it out ( Schmidt Heinrich , Haeckels Biogenetisches Grundgesetz und seine Gegner (Odenkirchen, 1902), 78–81). In Strasbourg, by contrast, Schwalbe regretted that Mehnert had rushed into speculation instead of completing his empirical work ( Schwalbe , “Mehnert” (ref. 68), 390–1); the essay on what Mehnert insisted on calling ‘Kainogenesis’ “almost led to my resignation from my position here” (see, also for his health and writing, Mehnert to Haeckel, 14 May 1898, Ernst-Haeckel-Haus Jena: Best. A, Abt. I). In 1898 Mehnert had nevertheless been hired as an associate professor of anatomy under Roux at Halle.

Schwalbe , “Variation” (ref. 65), 8.

NT 2, 2–3, 13. Fischel concluded conventionally that the range of variation declined as development proceeded, presumably as a result of correlation; see Fischel Alfred , “Über Variabilität und Wachsthum des embryonalen Körpers”, Morphologisches Jahrbuch, xxiv (1896), 369–404; and Keibel , “Das biogenetische Grundgesetz” (ref. 18), 764–8.

Keibel , “Normentafeln” (ref. 59). There was not even an honorarium; see von Lucius Annelise (Fischer-Verlag) to Friedrich Kopsch, 26 May 1952, Thüringisches Hauptstaatsarchiv Weimar: Archiv Verlagshaus Gustav Fischer Jena, Korrespondenzakte 1952, Kl—Lat. I am grateful for the limited access I was granted to this collection, but I was not permitted to consult several possibly-relevant files.

Keibel received support from the Prussian Academy of Sciences (NT 1, 10), and Otto Grosser and Julius Tandler from the Academy of Sciences in Vienna (NT 9, preface).

Taylor Ewing (NT 5) described the work as making his brain feel “rather stale”; see Taylor to Minot , 30 July 1904, Harvard Medical Library in the Francis A. Countway Library of Medicine, Boston, Mass.: H MS c21.2. On Minot's embryology, see Lewis Frederic T. , “Charles Sedgwick Minot”, The anatomical record, x (1916), 133–64.

Lillie F. R. , review of NT 11, The anatomical record, v (1911), 186.

Roy MacLeod drew attention to the relations between embryology and empire; see MacLeod Roy , “Embryology and empire: The Balfour Students and the quest for intermediate forms in the laboratory of the Pacific”, in idem and Rehbock Philip F. (ed.), Darwin's laboratory: Evolutionary theory and natural history in the Pacific (Honolulu, 1994), 140–65. I stress the intellectual and material transformations that scientists' appropriations worked, and their limits.

Clarke Adele E. , “Research materials and reproductive science in the United States, 1910–1940”, in Geison Gerald L. (ed.), Physiology in the American context, 1850–1940 (Bethesda, Md., 1987), 323–50, pp. 332–4.

NT 4, 2.

Keibel , “Entwicklungsgeschichte des Schweines” (ref. 56), 10–11; NT 6, preface; Fick , “Gedächtnisrede” (ref. 52), p. cxiii; NT 12, 1.

NT 9, preface, 1–2; http://www.fonyod.hu/, accessed 21 May 2004.

Eycleshymer Albert C. , “The habits of Necturus maculosus”, The American naturalist, xl (1906), 123–36, p. 132 (quote); NT 11, preface, 1; L[illie], review of NT 11 (ref. 76).

MacLeod , “Embryology and empire” (ref. 77).

Corner George W. , Ourselves unborn: An embryologist's essay on man (New Haven, 1944), 28.

Semon , Im australischen Busch (ref. 17); Kerr John Graham , A naturalist in the Gran Chaco (Cambridge, 1950); Hindle Edward , “John Graham Kerr, 1869–1957”, Biographical memoirs of Fellows of the Royal Society, iv (1958), 155–66.

Kerr J. Graham , “The development of Lepidosiren paradoxa: Part II. With a note upon the corresponding stages in the development of Protopterus annectens”, Quarterly journal of microscopical science, xlv (1901), 1–40 (p. 3 suggests that the Protopterus embryos for the normal plate also came from Budgett); Kerr J. Graham (ed.), The work of John Samuel Budgett, Balfour Student of the University of Cambridge. (Cambridge, 1907); Hall Brian K. , “John Samuel Budgett (1872–1904): In pursuit of Polypterus”, BioScience, li (2001), 399–407. Budgett's sacrifice for evolutionary embryology rivals the gathering of emperor-penguin eggs on the ‘winter journey’ of Scott's Antarctic expedition; see Raff Rudolf A. , The shape of life: Genes, development, and the evolution of animal form (Chicago, 1996), 1–4.

Hubrecht A. A. W. , “Studies from the Zoological Laboratory in the University of Utrecht: IV. Spolia nemoris”, Quarterly journal of microscopical science, xxxvi (1894), 77–125; Assheton Richard , “Dr Ambrosius Arnold Willem Hubrecht”, Proceedings of the Linnean Society of London, cxxvii (1915), 28–31; Keibel Franz , “A. A. W. Hubrecht: Ein Nachruf”, Anatomischer Anzeiger, xlviii (1915), 201–8; Faber J. , “Hubrecht, Ambrosius Arnold Willem”, in Dictionary of scientific biography, vi, 535–6; Bowler , Life's splendid drama (ref. 28), 181–3, 295–6.

Semon , Im australischen Busch (ref. 17); Kerr , Naturalist in the Gran Chaco (ref. 85), 173–5.

For complaints of ignorance, see Seidel Eduard , “19. Jahrhundert: Zur Vorgeschichte des Paragraphen 218”, in Jütte Robert (ed.), Geschichte der Abtreibung: Von der Antike bis zur Gegenwart (Munich, 1993), 120–39; for reframing, Hopwood , “Producing development” (ref. 8), 38–40; and idem, “Embryonen” (ref. 8), 240–1, 267–72.

NT 12, 1.

Kerr , Naturalist in the Gran Chaco (ref. 85), 175, 179, 180–1.

Budgett J. S. , “On the breeding-habits of some West-African fishes, with an account of the external features in development of Protopterus annectens, and a description of the larva of Polypterus lapradei”, Transactions of the Zoological Society of London, xvi (1901), 115–36, p. 120.

NT 7, 35. Hubrecht probably means a belief he reported earlier that the loris skeleton is “most efficacious in bringing about death and destruction among the unfortunate inhabitants of a house in front of which it has been buried overnight”. This generated “high demand among the wealthier natives who have family quarrels to settle, and I have known exorbitant prices, with which a collecting embryologist could not possibly compete, to be stealthily paid for one specimen, for this unfriendly though perhaps harmless purpose” ( Hubrecht , “Spolia nemoris” (ref. 87), 90–91).

Kerr , Naturalist in the Gran Chaco (ref. 85), 179.

Keibel , “Normentafeln” (ref. 59), 231, 233; idem, “Mitteilungen” (ref. 59), 594.

The two prewar exceptions are Semon, who used a cheaper half-tone process because his figures had already been published in excellent lithographs (NT 3, iii), and Grosser and Tandler, who probably favoured photomechanical reproduction because they began with photographs that an artist retouched and made more detailed (NT 9, 2–3). Keibel , finding the retouched photograms “somewhat dull”, preferred lithographs, but left the decision to Tandler and Fischer; see Tandler Keibel to , 19 March 1906, and Fischer to Keibel, 28 August 1908, Institut für Geschichte der Medizin der Medizinischen Universität Wien: HS 4508/3 and 4066/4. Peter had drawings done after photographs, but mostly not directly onto them (NT 4, 3–4).

NT 3, 2; NT 4, 3.

Daston Galison , “Image of objectivity” (ref. 4); Hopwood , “Producing development” (ref. 8), 73–74.

Kerr , “Development of Lepidosiren” (ref. 86), 36–37; NT 10.

NT 5, 1; NT 12, 1. On reconstructions, see also Lewis Frederic T. , review of NT 8, Science, xxix (1909), 939–40. To get to grips with complex structures wax models were sometimes reconstructed from the serial sections and occasionally even depicted on normal plates (NT 1, 11, pl. II–III), an indication of the centrality of modelling in vertebrate embryology, on which see Hopwood , “‘Giving body’” (ref. 5); and idem, Embryos in wax (ref. 8).

NT 12, 70.

NT 8; Hopwood , “Producing development” (ref. 8), 74–76.

NT 12, 70–78; Balfour , Development of elasmobranch fishes (ref. 10), 71–80, pl. VI-VII.

Gould , Ontogeny and phylogeny (ref. 11), 174.

E.g., NT 7, 28–34, 48–61; NT 8, 152–62; NT 9, 43. Bruno Henneberg, believing his penultimate volume likely to be the last, attempted to summarize the relative rates at which different organs developed among the amniotes in the series, and concluded that there was considerable agreement (NT 15, 110–60).

Gould , Ontogeny and phylogeny (ref. 11), 174; Bowler , Life's splendid drama (ref. 28). Keibel's own position changed little; see Keibel Franz , “Die Entwicklungsgeschichte der Wirbeltiere”, in Hinneberg Paul (ed.), Die Kultur der Gegenwart: Ihre Entwicklung und ihre Ziele, part iii, section iv, vol. ii: Zellen- und Gewebelehre, Morphologie und Entwicklungsgeschichte, part 2: Hertwig O. (ed.), Zoologischer Teil (Leipzig, 1913), 333–98.

Pfühl Wilhelm , “Karl Petert”, Anatomischer Anzeiger, cii (1955), 224–44.

Keibel , “Das biogenetische Grundgesetz” (ref. 18), 758.

Kerr J. Graham , Text-book of embryology, ii: Vertebrata with the exception of Mammalia (London, 1919), 575. See also Anon., review of NT 1, Revue scientifique, 1898; C. A. K., review of NT 2, Journal of applied microscopy and laboratory methods, both consulted as clippings in the Fischer Archiv (ref. 73), Rezensionsarchiv, Karton 40; and Waldeyer Wilhelm , “Anatomie: X. Lehr- und Handbücher”, Ergebnisse der Anatomie und Entwicklungsgeschichte, xii (1903), 652–742, p. 715, judging NT 1–3 “indispensable for all further investigations”.

For the standard run, see Von Lucius to Kopsch , 26 May 1952 (ref. 73). The prices (from the wrappers of NT 10–12) range from 7.50 marks for NT 9 to 36 marks for NT 8; as a comparison, Oscar Hertwig's textbook cost 13 and his 6-volume handbook 135 marks.

On Tandler, see Sablik Karl , Julius Tandler, Mediziner und Sozialreformer: Eine Biographie (Vienna, 1983); and for the suggestion that Abraham, the leading figure in German psychoanalysis, “never lost” the interest in embryology gained in work on NT 2 and his dissertation on budgerigar development, Abraham Hilda C. , “Karl Abraham: An unfinished biography”, International review of psychoanalysis, i (1974), 17–72, p. 22; see also Scammon Richard E. Calkins Leroy A. , The development and growth of the external dimensions of the human body in the fætal period (Minneapolis, 1929).

Gursch , Die Illustrationen Ernst Haeckels (ref. 43), 84–136; Daum Andreas W. , Wissenschaftspopularisierung im 19. Jahrhundert: Bürgerliche Kultur, naturwissenschaftliche Bildung und die deutsche Öffentlichkeit, 1848–1914 (Munich, 1998), 210–35; Keibel Franz , “Haeckel und Brass”, Deutsche medizinische Wochenschrift, xxxv (1909), no. ii, 350–1.

Mall Franklin P. , review of NT 8, The anatomical record, ii (1908), 368–71, p. 371; see also Lewis , review of NT 8 (ref. 100). Keibel had the book sent to Mall and asked for a review; see Mall Keibel to , 23 July 1908, Carnegie Department Papers (ref. 61), record group I, series 1, box 10, folder 34.

Haeckel Ernst , “Die Grenzen der Naturwissenschaft”, Deutsche medizinische Wochenschrift, xxxvi (1910), no. ii, 1855–7, p. 1857 (quotes); idem, Anthropogenie, 5th edn (ref. 70), i, 380; Schmidt Heinrich , Haeckels Embryonenbilder: Dokumente zum Kampf um die Weltanschauung in der Gegenwart (Frankfurt am Main, 1909), 81–87.

On Keibel's career, see especially Nauck Keibel (ref. 52), 18–25; and on the anatomical job market, Nyhart , Biology takes form (ref. 11), 278–305.

His Wilhelm , “Antrag der Königlich Sächsischen Gesellschaft der Wissenschaften auf Bestellung einer Fachcommission für menschliche und thierische Entwickelungsgeschichte und für Anatomie des Gehirnes, vorgelegt der Generalversammlung der Internationalen Association der Akademien am 16. April 1901 zu Paris”, Berichte über die Verhandlungen der Königlich Sächsischen Gesellschaft der Wissenschaften zu Leipzig, Mathematisch-physische Classe, liii (1901), 77–82. The Association (on which see Greenaway Frank , Science international: A history of the International Council of Scientific Unions (Cambridge, 1996), 6–18) decided to constitute a special commission for brain anatomy, but for the moment to leave the smaller and more divided field of embryology to the subject societies; the Anatomische Gesellschaft created a commission of Wilhelm Waldeyer, Carl Rabl, Keibel and His (His, “Wissenschaftliche Centralanstalten” (ref. 59), 414).

Nieuwkoop P. D. , “‘L'Institut International d'Embryologie’ (1911–1961)”, General embryological information service, ix (1961), 265–9. For a record of the first two meetings, see [ Keibel Franz Assheton Richard Hubrecht A. A. W. , “Institut International d'Embryologie: Session de 1912”, Bibliographie anatomique, xxii, no. 3, offprint numbered 1–12, in Carnegie Department Papers (ref. 61), record group I, series 1, box 9, folder 27. The Institut was also interested in supporting the International Commission on Embryological Nomenclature. Of the 11 men at the founding meeting, eight were among Keibel's actual or prospective authors. Nieuwkoop's and other accounts name Hubrecht, Keibel and the British embryologist Richard Assheton as the initiators of the Institut, but Bonnet was in fact the third signatory of the call, was (unlike Assheton) at the founding meeting, and was even the first president. He may have been written out of the history for unclear reasons associated with the resignation of his membership by the following year.

Nieuwkoop , “‘L'Institut International’” (ref. 117), 265–6; Faasse Patricia Faber Job Narraway Jenny , “A brief history of the Hubrecht Laboratory”, International journal of developmental biology, xliii (1999), 583–90; Richardson Michael K. Narraway Jennifer , “A treasure house of comparative embryology”, ibid., 591–602. Faasse Patricia , Zuiver om de wetenschap: De Akademie en haar levenswetenschappelijke instituten (Amsterdam, 1999), 27–30, makes clear that the idea of a central embryological collection went back to His's proposals.

On the early years of the Carnegie Department, see Sabin Florence Rena , Franklin Paine Mall: The story of a mind (Baltimore, 1934); O'Rahilly , “One hundred years” (ref. 10); and Morgan Lynn M. , “Properly disposed of: A history of embryo disposal and the changing claims on fetal remains”, Medical anthropology, xxi (2002), 247–74. For Mall's extensive discussions with Keibel over founding what became the Contributions to embryology, see their correspondence in Carnegie Department Papers (ref. 61), record group 1, series 1, box 10; and on the money, specifically Keibel to Mall, 23 February, 10 April and 18 June 1914, ibid., folder 37.

Peter , “Keibel” (ref. 52), 202–4; Fick , “Gedächtnisrede” (ref. 52), pp. cix–cxi; Weindling , Darwinism (ref. 26), 250.

Keibel to Fischer , 10 and 15 March 1922, Fischer Archiv (ref. 73), Korrespondenzakten.

Keibel to Fischer , 24 January 1929, Ibid.

Fischer to Keibel , 9 December 1922, ibid.; Keibel to Streeter George L. , 13 December 1922 and 25 March 1924, Carnegie Department Papers (ref. 61), record group I, series 2, box 26, folder 24.

Glaesner had also been an assistant at the Berlin oceanography museum; see Nyhart Lynn K. , “Science, art, and authenticity in natural history displays”, in de Chadarevian Hopwood (ed.), Models (ref. 8), 307–35, pp. 323–7.

de Lange Dan Jr Nierstrasz H. F. , Tabellarische Uebersicht der Entwicklung von Tupaia javanica Horsf. (Utrecht, 1932); Huisman F. J. de Lange Dan , Tabellarische Uebersicht der Entwicklung von Manis javanica Desm. (Utrecht, 1937). These volumes are smaller than Keibel's. On the revival of the I. I. d'E., negotiations over and promotion of the plates, see the German, English and French introduction to the de Lange and Nierstrasz plate; and correspondence of the I. I. d'E. with Warren H. Lewis, American Philosophical Society, Philadelphia: B L586; and with Streeter, 1931–8, Carnegie Department Papers (ref. 61), record group I, series 2, box 24, folders 25–26.

For American demand for a rat Normentafel, see Streeter to Keibel , 5 March, 28 April and 11 June, and Keibel to Streeter, 25 March and 1 April 1924, Carnegie Department Papers (ref. 61), record group I, series 2, box 26, folder 24; on the much delayed production of the plate, which unusually includes many figures of sections, see correspondence between Bruno Henneberg and Fischer, Fischer Archiv (ref. 73), Korrespondenzakten 1937, Hart-Hern; and for its status as Henneberg's “main scientific work”, see Duncker Hans-Rainer , “Bruno Henneberg (1867–1941) /Anatom”, in Gundel Hans Georg Moraw Peter Press Volker (ed.). Gießener Gelehrte in der ersten Hälfte des 20. Jahrhunderts (2 parts, Marburg, 1982), Part 1, 378–86, p. 384.

Richardson Michael K. , “Heterochrony and the phylotypic period”, Developmental biology, clxxii (1995), 412–21; Bininda-Emonds Olaf R. P. , Jeffery Jonathan E. Coates Michael I. Richardson Michael K. , “From Haeckel to event-pairing: The evolution of developmental sequences”, Theory in biosciences, cxxi (2002), 297–320, pp. 301–3.

Willier Benjamin H. Weiss Paul A. Hamburger Viktor , Analysis of development (Philadelphia, 1955), p. v.

Oppenheimer Jane M. , “Problems, concepts and their history”, ibid., 1–24, p. 22; idem, “Methods and techniques”, ibid., 25–38.

Idem, “The normal stages of Fundulus heteroclitus”, The anatomical record, lxviii (1937), 1–8; idem, “Fifty years of Fundulus”, The quarterly review of biology, liv (1979), 385–95, p. 386. This was in a Yale tradition, which, as I am about to describe, went back (through John S. Nicholas) to Harrison.

Hamburger Viktor Hamilton Howard L. , “A series of normal stages in the development of the chick embryo”, Journal of morphology, lxxxviii (1951), 49–92, reprinted in Developmental dynamics, cxcv (1992), 231–72; Sanes Joshua R. , “On the republication of the Hamburger-Hamilton stage series”, ibid., 229–30.

Pickstone John V. , “Museological science? The place of the analytical/comparative in nineteenth-century science, technology and medicine”, History of science, xxxii (1994), 111–38. On the literary control of variability in the laboratory, see Köhler , Lords of the fly (ref. 2), 71–77.

Peter Karl , “Der Grad der Beschleunigung tierischer Entwicklung durch erhöhte Temperatur”, Archiv für Entwicklungsmechanik der Organismen, xx (1906), 130–54, p. 134.

Harrison Ross G. , “Experiments on the development of the fore limb of Amblystoma, a self-differentiating equipotential system”, The journal of experimental zoology, xxv (1918), 413–59, p. 417; idem, Organization and development of the embryo, ed. by Wilens Sally (New Haven, 1969), 45. On Harrison, see especially Maienschein, Transforming traditions (ref. 11), 261–89.

Harrison Ross G. , “The development of the balancer in Amblystoma, studied by the method of transplantation and in relation to the connective-tissue problem”, The journal of experimental zoology, xli (1925), 349–427, pp. 359, 361.

Harrison to the Chicago anatomist, Herrick C. Judson , 26 March 1937, Yale University Library, Manuscripts and Archives: Ross Granville Harrison Papers (MS 263), series I, box 12, folder 884.

Schwind Joseph L. Twitty Victor C. to Harrison , 3 October 1927, ibid., series III, box 38, folder 257.

Harrison , Organization and development (ref. 134), 44–66.

Quote from Lehman H. E. , “A memorial in embryology”, Science, clxviii (1970), 724–5; see also Hamburger Viktor , review of Harrison, Organization and development (ref. 134), American scientist, lviii (1970), 321–2.

Pollister Arthur W. Moore John A. , “Tables for the normal development of Rana sylvatica”, The anatomical record, lxviii (1937), 489–96, p. 489.

For a distribution list, see Harrison Papers (ref. 136), series III, box 38, folder 258.

Swett Francis H. to Harrison , 21 November 1928, Ibid.

Hamburger Viktor , A manual of experimental embryology (Chicago, 1942), 196–204. Rugh Compare Roberts , Experimental embryology: A manual of techniques and procedures, rev. edn (Minneapolis, 1948), 56–101; Rugh to Harrison , 25 February and 28 March 1941, Harrison Papers (ref. 136), series I, box 23, folder 1727. To my knowledge, the stage descriptions were first published in Harrison's book.

Hamburger Viktor , “Über den Einfluss des Nervensystems auf die Entwicklung der Extremitäten von Rana fusca”, Wilhelm Roux' Archiv für Entwicklungsmechanik der Organismen, cv (1925), 149–201, pp. 157–8; idem, “Afterword: The stage series of the chick embryo”, Developmental dynamics, cxcv (1992), 273–5, p. 273. Hamburger's own account suggests he was inspired by Harrison's work, and the Yale zoologist may have been more important to him than the Freiburg anatomical tradition, but he appears to have obtained photographs from Harrison only in 1929 (Hamburger to Harrison, 10 March 1929, Harrison Papers (ref. 136), series I, box 11, folder 828). On Hamburger, see his The heritage of experimental embryology: Hans Spemann and the organizer (New York, 1988); and “Viktor Hamburger virtual exhibit”, http://library.wustl.edu/units/biology/vh/, accessed 24 May 2004.

Waelsch Salome G. , “The causal analysis of development in the past half century: A personal history”, Development, 1992, supplement, 1–5, p. 1 (quotes); Glücksohn Salome , “Äussere Entwicklung der Extremitäten und Stadieneinteilung der Larvenperiode von Triton taeniatus Leyd. und Triton cristatus Laur.”, Wilhelm Roux' Archiv für Entwicklungsmechanik der Organismen, cxxv (1931), 341–405; Hamburger , “Afterword” (ref. 144), 274; Fäßler Peter E. , Hans Spemann 1869–1941: Experimentelle Forschung im Spannungsfeld von Empirie und Theorie. Ein Beitrag zur Geschichte der Entwicklungsphysiologie zu Beginn des 20. Jahrhunderts (Berlin, 1997), 70–72 (on Spemann's treatment of female doctoral candidates). Hamburger sought Harrison's permission for Waelsch to refer to his stages; see Hamburger to Harrison , 21 November 1930 (quote), and Harrison to Hamburger , 27 June 1931, Harrison Papers (ref. 136), series I, box 11, folder 828.

Hamilton Howard L. , Lillie's development of the chick: An introduction to embryology, 3rd edn (New York, 1952), 74–91; Hamburger Hamilton , “Normal stages” (ref. 131); Hamburger , “Afterword” (ref. 144).

For critical remarks on NT 2, see Hamburger Hamilton , “Normal stages” (ref. 131), 50; and on NT 4, Pasteels Jean J. , “Une table analytique du développement des reptiles: 1. Stades de gastrulation chez les Chéloniens et les Lacertiliens”, Annales de la Société Royale Zoologique de Belgique, lxxxvii (1957), 217–41, pp. 217–18. For Glücksohn's explanation why she followed Harrison's stages rather than Glaesner's (NT 14), see her “Äussere Entwicklung” (ref. 145), 353. See also Hamburger to Harrison , 21 November 1930, Harrison Papers (ref. 136), series I, box 11, folder 828.

See Hamburger Hamilton , “Normal stages” (ref. 131), 50, for one of many claims that earlier controversies could have been avoided had proper stages been used.

Hamburger , “Afterword” (ref. 144), 275.

Hamburger Hamilton , “Normal stages” (ref. 131), 52.

Kopsch Friedrich , Die Entwicklung des braunen Grasfrosches Rana fusca Roesel, dargestellt in der Art der Normentafeln zur Entwicklungsgeschichte der Wirbeltiere (Stuttgart, 1952), p. vi. Quote from Keibel to Fischer Gustav , [1922], Fischer Archiv (ref. 73), Korrespondenzakte 1922, Kla–Kly. Kopsch switched to a Stuttgart publisher because obtaining permission to publish the work with Fischer in Jena threatened to take too long. On Kopsch, see Richter W. , “Friedrich Kopsch und sein Beitrag zur Entwicklung der Berliner Anatomie im 19. und 20. Jahrhundert”, Charité-Annalen, n. s., viii (1988), 237–42.

Nieuwkoop Faber (eds), Normal table (ref. 1), 1; Gurdon Hopwood , “Introduction of Xenopus” (ref. 1). Funding was obtained from various Dutch organizations and (for publication) from the International Council of Scientific Unions, successor to the International Association of Academies ( Nieuwkoop Faber (ed.), Normal table (ref. 1), 3).

Nieuwkoop Faber (ed.), Normal table (ref. 1); additional information from Faber to the author, 14 July 1999.

Witschi Emil , “Proposals for an international agreement on normal stages in vertebrate embryology”, XIV International Congress of Zoology, Copenhagen, 5.–12. August 1953, proceedings (Copenhagen, 1956), 260–2. Ronan O'Rahilly of the Carnegie Department found the idea “attractive”; see O'Rahilly , Developmental stages (ref. 10), 7–8; and idem and Müller , Developmental stages (ref. 10), 8.

Oppenheimer Jane M. , “The growth and development of developmental biology”, in Locke Michael (ed.), Major problems in developmental biology (New York, 1966), 1–27; Keller Evelyn Fox , Refiguring life: Metaphors of twentieth-century biology (New York, 1995); Hopwood , “Embryology” (ref. 3).

Gurdon Hopwood , “Introduction of Xenopus” (ref. 1); for ‘literary technology’, see Shapin Steven , “Pump and circumstance: Robert Boyle's literary technology”, Social studies of science, xiv (1984), 481–520.

In the late 1980s and 1990s, many laboratories switched to the zebrafish and consulted Kimmel Charles B. , Ballard William W. , Kimmel Seth R. , Ullmann Bonnie and Schilling Thomas F. , “Stages of embryonic development of the zebrafish”, Developmental dynamics, cciii (1995), 253–310.

Griesemer James Yamashita Grant , “Managing time in model systems: Illustrations from evolutionary biology”, unpublished paper. Other aspects of the history of normal plates could be analysed in time management terms, such as Keibel's enlisting collaborators and the use of Xenopus to overcome seasonality.

O'Rahilly , “One hundred years” (ref. 10), 93.

On the Carnegie Department at mid-century and since, see Corner , Ourselves unborn (ref. 84); McLaughlin Loretta , The pill, John Rock, and the Church: The biography of a revolution (Boston, 1982), 58–71; Brown Donald D. , “The Department of Embryology of the Carnegie Institution of Washington”, BioEssays, vi (1987), 92–96; and http://nmhm.washingtondc.museum/collections/hdac/index.htm, accessed 3 June 2004.

Streeter George L. , “Department of Embryology”, Carnegie Institution of Washington year book, xxi (1922), 76–92, p. 76.

Idem, “Developmental horizons in human embryos: Description of age group XI, 13 to 20 somites, and age group XII, 21 to 29 somites”, Contributions to embryology, xxx (1942), 211–45, pp. 213–14; Mall Franklin P. , “On stages in the development of human embryos from 2 to 25 mm long”, Anatomischer Anzeiger, xlvi (1914), 78–84. On Streeter, see Corner George W. , “George Linius Streeter, 1873–1948”, Biographical memoirs of the National Academy of Sciences, xxviii (1954), 260–87.

Streeter George L. , “Developmental horizons in human embryos: Description of age groups XV, XVI, XVII, and XVIII, being the third issue of a survey of the Carnegie Collection”, Contributions to embryology, xxxii (1948), 133–203, p. 135.

Idem, “Developmental horizons in human embryos: Description of age groups XIX, XX, XXI, XXII, and XXIII, being the fifth issue of a survey of the Carnegie Collection”, ibid., xxxiv (1951), 165–96, p. 169.

Idem, “Third issue of a survey” (ref. 163), 137.

Idem, “Archetypes and symbolism”, Science, lxv (1927), 405–12.

O'Rahilly , Developmental stages (ref. 10), 4; idem and Müller , Developmental stages (ref. 10).

O'Rahilly , Developmental stages (ref. 10), 7–8. Carnegie stages are still in general use, but there are signs that they no longer meet all the demands of a human embryology that has been transformed in the last few decades ( Barbet Patrick , “Une approche historique de l'étude des stades du développement de l'embryon humain”, talk presented at workshop, “L'embryon humain face au temps de l'histoire, face au temps de la vie”, Fribourg, Switzerland, 29 October 2004).

Galison , “Judgment against objectivity” (ref. 4).

Ibid., 354.; Daston , “Objectivity versus truth” (ref. 4).

Quotes from Corner, preface to Streeter's Developmental horizons (ref. 10).

During the First World War Mall obtained embryos of various species for Keibel; their letters illustrate the utility of normal plates in communicating with collectors. Keibel sometimes used the plates to describe stages he needed (Keibel to Mall, 7 November 1915 and 14 March 1916, Carnegie Department Papers (ref. 61), record group 1, series l, box 10, folder 40), but Mall “could not get any standard plates on the turtle, so it was impossible for me to place the order with the collector in a more definite way” (Mall to Keibel, 24 January 1916, ibid.).